Eastern Carpathians, Ukraine)

Plant and Fungal Systematics 64(1): 53–64, 2019 ISSN 2544-7459 (print) DOI: 10.2478/pfs-2019-0007 ISSN 2657-5000 (online)

Dedicated to the late Professor Jadwiga Siemińska

Green and charophytic algae of the high-mountain Nesamovyte and Brebeneskul lakes (Eastern Carpathians, Ukraine)

Petro Tsarenko1*, Konrad Wołowski2, Joanna Lenarczyk2, Olena Bilous3 & Halyna Lilitska1

Abstract. A study of green and charophytic algae diversity in two high-mountain lakes in Article info the Eastern Carpathians (Ukraine) identified 99 species (109 taxa at species and intraspecific Received: 5 Feb. 2019 rank) in 35 genera from different ecotopes of the studied lakes. Algal species composition was Revision received: 19 Apr. 2019 characterized, and the ecological parameters of the lakes were determined from monitoring Accepted: 10 May 2019 data recorded over the last century. Environmental analyses using bioindication methods Published: 30 Jul. 2019 based only on data on the composition of green and charophytic algae confirmed that the Associate Editor environmental inferences were accurate. Degradation of the Nesamovyte and Brebeneskul Adam Flakus lake ecosystems, as compared with their earlier states, was noted. Key words: algae, diversity, Eastern Carpathians, mountains lakes, bioindication, ecosystems

Introduction

Algae are an important component of mountain ecosys- Buczkó et al. 2009; Kot 2009; Cărăus 2012; Lenarczyk tems, where they form a particular spectrum of taxonomic 2012; Lenarczyk & Tsarenko 2013). groups, with montane forms as well as rare and distinctive Data on algal diversity in the Eastern (Ukrainian) Car- species. Algae exhibit geographical, zonal and stratigraphic pathians, and especially on their individual taxonomic specificity in their species composition and in their asso- groups and on the mountain ranges, are scarce and incom- ciation with types of waterbodies and biotopes, and are plete (Vodopian 1981; Palamar-Mordvintseva et al. 1992; considered to be indicators of climatic changes (Nauwerk Tsarenko et al. 1997, 1998, 2009, 2014, 2017; Tsarenko 1966; Ettl 1968; Wasser 1989; Lampert & Sommer 1997; & Parchuk 1998; Tsarenko 2000; Tsarenko & Wasser 2000; Kamenik et al. 2000; Nedbalova et al. 2006; Barinova et al. Tsarenko & Palamar-Mordvintseva 2014, 2016; Myki- 2006, 2019; Cantonati & Spitale 2009; Bąk et al. 2012). Of tchak 2014; Tsarenko & Lilitska 2016; Bilous & Tsarenko particular indicator value are diatoms and green and cha- 2018). This applies primarily to the waterbodies of the rophyte algae, which often are subdominants or dominants high-mountain ecosystems of this region and specifically in the waterbodies of these ecosystems (Kawecka 1970; to the largest subalpine lakes of the Chornohora Mts of Kopaček et al. 2004; Buczko & Wojtal 2007; Kawecka the Eastern Carpathians in Ukraine (Tsarenko et al. 2014). & Galas 2003; Kawecka & Robinson 2008; Robinson et al. Published algofloristic information for this area comes 2010; Burchardt 2014; Wojtan et al. 2014; Khuram et al. from the 20th century (Wołoszynska 1920; Asaul 1969; 2019). In terms of composition, taxonomic structure and Palamar-Mordvintseva 1978a, b, 1982) and needs updat- distribution, algal diversity is high in the mountain water- ing; the current state of algal diversity and species com- bodies of the Carpathian region (Szklarczyk-Gazdowa 1960; position needs to be documented. Recently we obtained Sieminska 1967; Hindák & Kováčik 1993; Lukavsky 1994; a general idea of the diversity and indicator value of algae Wojtal & Galas 1994; Fott 1999; Juriš & Kováčik 1987; in the lentic system of Chornohora (Tsarenko et al. 2014, 2015, 2017, 2018). The diatom species composition of Nesamovyte and Brebeneskul lakes, and also Маricheyka 1 Department of Phycology, Lichenology and Briology, M.G. Kholodny, Lake from the forest zone of the Carpathians, was studied National Academy of Science of Ukraine, 2 Tereschenkivska St., 01004, Kyiv, Ukraine in detail (Kryvosheya & Tsarenko 2018; Tsarenko et al. 2 Department of Phycology, W.Szafer Institute of Botany, Polish 2016). In the present study we addressed the present-day Academy of Sciences, ul. Lubicz 46, 31-512 Krakow, Poland species composition of “green” algal phyla (Chlorophyta, 3 Department of Ecology of Aquatic Plants and Toxicology, Institute of Charophyta) in the highest lakes of Chornohora (Nesa- Hydrobiology, National Academy of Science of Ukraine, 12 Geroiv Stalingrada Ave., 04210, Кyiv, Ukraine movyte, Brebeneskul), analysed monitoring data for the * Corresponding author e-mail: [email protected] last century, and determined the ecological state of these

This work is licensed under the Creative Commons BY-NC-ND 4.0 License 54 Plant and Fungal Systematics 64(1): 53–64, 2019

lakes on the basis of algal indicator species occurring Table 1. Morphometry and basic parameters of the studied lakes (orig- under recent environmental conditions1. inal measurements)

Location Nesamovyte Lake Brebeneskul Lake Materials and methods 48°07ʹ36.6ʺN 48°06ʹʹ06.0ʺN Coordinates 24°32ʹ26.4ʺE 24°33ʹ44.2ʺE The investigated lakes are located in the highest part of the Elevation, m a.s.l. 1748 1793 Ukrainian Carpathians (Fig. 1), the Polonyno-Chornohora Area, ha 0.3 0.6 geomorphological area (Tsys 1968), which is part of the Max. depth, m ~2.0 2.4 Carpathian-Danube algofloristic sub-provinces according Water temp., °C 18.3 (16.2) 18.4 (–12.6) to the algofloristic zonation of Ukraine (Palamar-Mord- pH 6.2–6.4 7.1 vintseva & Tsarenko 2015). Conductivity, μS/cm 8.2–9.8 26.74 The material for the study (16 samples of net plankton, O2, mg/l 10.7 6.32 the periphyton communities of algae on higher aquatic plants, and squeezes from moss from marshland) was habitat preference (B – benthic; P-B – plankto-benthic; collected along the perimeter of the lakes (more along S – soil; Ep – epiphyte; P – planktonic), current and the southern parts) in the summers of 2013–2017 (July– oxygenation (аer – aerophyles; St-str – inhabitants of August 2013–15, August 2016–2017). Nesamovyte Lake low-flow, moderately oxygenated water; ae – aerophytes; is in a glacial cirque on the eastern slope of Mount Turkul st – inhabitants of standing water with low oxygenation), (Chornohora Ridge, Eastern Carpathians) at 1750 m a.s.l. pH (Hustedt 1957) (ind – indifferent; acf – acidophile), The lake covers ~0.3 hectares (88 × 45 m) and is ~2 m salinity (Hustedt 1957) (i – indifferent; hb – halophobe; deep. It is characterized by atmospheric supply of water oh – oligohalobe), trophic state (Van Dam et al. 1994) and food, with a sandy/muddy bottom and a long freez- (m – mesotraphentic; me – meso-eutraphentic; o-m – ing period. The water in the lake is now β-mesosaprobic oligo- to mesotraphentic; e – eutraphentic; ot – oligotrap- (Mikitschak & Kokish 2014). hentic), and organic pollution (Sladecek 1973; Barinova Brebeneskul Lake is one of Ukraine’s highest moun- et al. 2006, 2019) (x – xenosaprobe; x-o – xeno-oligosap- tain lakes. It is in a glacial cirque on the southwestern robe; o-x –oligo-xenosaprobe; x-b – xeno-betamesosap- slope of Chornohora Ridge at 1793 m a.s.l. in a depres- robe; o – oligosaprobe; o-b – oligo-betamesosaprobe; sion between Mt. Brebeneskul (2035 m a.s.l.) and Mt. b-o – beta-oligosaprobe; o-a – oligo-alphamesosaprobe; Gutin-Tomnatek (2016 m a.s.l.) in the Chornohora area of b – betamesosaprobe; b-a – beta-alphamesosaprobe; a – the Carpathian Biosphere Reserve (Mykitchak & Kokish alpha-mesosaprobe). Organic pollution was calculated 2014). The lake covers ~0.6 ha (146 × 67 m) and is up to based on the values of the saprobity indices and indicator 2.4 m deep. The lake is not drained but undergoes weak groups, and, accordingly, water quality classes. The above filtration through the ridge from the eastern side, and is indicators were used to assign the water to the following supplemented by atmospheric precipitation and ground- quality classes: I – 0–0.5 (x, x-o); II – 0.6–1.5 (o-x, x-b, water. The bottom is stony silt, and the weakly miner- o, o-b); III – (b-o, o-a, b, b-a); and IV – (a) (Belous et al. alized water is oligosaprobic (Mykitchak 2014). During 2013; Bilous et al. 2014, 2016; Barinova et al. 2019). the study period the water parameters were as follows: Valid names are given according to AlgaeBase (Guiry 18.4–12.6°C (upper to lower layer), pH 7.1, specific con- and Guiry 2019). The taxonomic list was compiled in ductivity 26.74 μS/cm, turbidity 3.91 NTU (nephelometric accordance with the system used in Tsarenko et al. turbidity units) and oxygen saturation 6.32 mg/L. (2011, 2014). This study is based on living algal material from the plankton (accumulation cultures, with addition of liquid and agar Bold Basal Medium; Andersen 2005) and on samples fixed with 4% formaldehyde solution. In addition to the algological samples we collected in 2013–2017, we studied material from the Algoteca of the M.G. Kholodny Institute of Botany, National Academy of Science of Ukraine (AKW, accessions NN 16855–16898, samples from 1967 collected by Z.I. Asaul). The algae were observed and identified by light microscopy (Ergaval Carl Zeiss, Olympus BX-53). Our ecological analyses of the two lakes is based on historical data for 1920 (only Nesamovyte Lake) and 1967 (both lakes), and recent data from 2013–2016. The following ecological characteristics were used:

1 This work was done under a Polish–Ukrainian agreement on scientific cooperation between the Polish Academy of Sciences (W. Szafer Institute of Botany, Poland) and the National Academy of Science of Figure 1. Location of Nesamovyte and Brebeneskul lakes (IF – Iva- Ukraine (M.G. Kholodny Institute of Botany, Ukraine). no-Frankivsk Region; ZK – Zakarpatia Region). P. Tsarenko et al.: Green and charophytic algae of the high-mountain lakes in the Eastern Carpathians 55

To compare the species composition of the lakes, only noted the presence of more than 100 Desmidiales we used Venny 2.1.0 software to construct Euler circles. taxa in 14 lakes of the region, without specifying the Species occurrence in the samples was estimated on the lakes, and mentioning several species of that group as Starmach scale (Wasser 1989). Ecological features of the well as coccoid green algae in the two lakes we studied species are presented according to Barinova et al. (2006, (Palamar-Mordvintseva 1978a, b). That author noted the 2015, 2019) and Bilous et al. (2016). distinctiveness of the algal species composition of the Chornohora region and characterized the lakes as a dif- Results and discussion ferent type of waterbody, and also determined the main factors contributing to the diversity and distribution of The material from 100 years of investigation of the two Desmidiales in the studied water bodies. lakes contained 99 species of green and charophytic algae The algological samples housed in AKW, taken from (109 taxa at species and intraspecific rank) belonging these lakes in the 1960s, archival materials from Prof. to four classes (Trebouxiophyceae, Chlorophyceae and G.M. Palamar-Morvintseva’s studies, and the results of Oedogoniophyceae from Chlorophyta; Zygnematophy- our research in 2013–2017 allowed us to track the species ceae from Charophyta), eight orders, 16 families and 35 diversity of the green and charophytic algae of Nesam- genera (Table 2). Charophytic algae account for most of ovyte and Brebeneskul lakes over time. Our research con- the algal diversity (84.4%). The families Desmidiaceae firmed the diversity of Desmidiales in Nesamovyte Lake (75.2%) of the charophytic algae and Scenedesmaceae and its marsh (Table 2). A comparison of the algal species (6.5%) of the green algae together formed ~82% of the composition of this lake at 50-year intervals (1910–1967– species composition of the algae. The richest genera were 2017) indicates a decrease in the species diversity of this Staurastrum (22 species, 25 intraspecific; 22.9%), Cos- group. There were 41 species (43 intraspecific) in the early marium (17 species, 15.6%), Euastrum (13 species, 17 20th century (Wołoszynska 1920), 29 (31 intraspecific) intraspecific; 15.6%), Closterium (6 species, 5.5%), Tet- in the 1960s (Palamar-Mordvintseva 1978a, b), and only memorus (2 species, 4 intraspecific; 3.7%),Actinotaenium 20 species recently. In the latter period, rare species of and Staurodesmus (3 species, 2.8% each), Pseudopedias- the genera Hyalotheca, Euastrum and Tetmemorus were trum (2 species, 3 intraspecific; 2.8%), andGonatozygon observed; they are peculiar to waterbodies of mountain and Penium (2 species, 1.8% each). Twenty-one genera regions. However, taxa of the genera Actinotaenium, were represented by only one species each. Cylindrocystis, Micrasterias, Netrium, Teilingia and About 85% of the species composition were charo- Coelastrum, noted in material from 1910, were not con- phytic algae (Zygnematophyceae), which develop pri- firmed, representatives of generaPenium , Sphaerozosma, marily in higher aquatic plant associations (58%). The Spirotaenia and Tortitaenia, found in material from 1967, share of this group in other biotopes is much lower: were not noted in 2017. Along with this, species of the ~20% in the benthos and ~22% in the plankton. We genera Mougeotia, Spirogyra, Zygnema, Oedogonium, found similar shares of ~85% for Chlorophyta in the Botryococcus, Chlamydomonas, Mucidosphaerium, Dic- water column of the two lakes, and small shares (7.5% tyosphaerium, Mychonastes and Westella appeared in each) in the benthos and in aquatic plant associations. 2017 for the first time. The presence of representatives Such an uneven biotopic distribution of algae is due to of the latter genera apparently indicates degradation of the ecological requirements and habit of the species and the oligotrophic waterbody, its transformation to a mes- taxonomic groups. Most of the species diversity was otrophic state, and its colonization by common species contributed by the algae of Nesamovyte Lake (83 species, having a wide ecological amplitude. 92 intraspecific; 84.4%). There was also a change in the quantitatively dom- The first information about the diversity of algae in inant species: at the beginning of the 20th century the Chornohora lakes, including this lake, was presented (Wołoszynska 1920) the dominants were Cylindrocystis by Woloszynska (1920), based on material from the early brebissonii, Actinotaenium cucurbita, Cosmarium 20th century (collected by R. Raciborski in 1910 and staurastriforme, C. venustum var. excavatum, Euastrum T. Wilezyński in 1914). Wołoszynska studied and ana- insigne, E. humerosum var. humerosum and var. subinter- lyzed the algal diversity of Nesamovyte Lake in detail medium, E. didelta and Staurastrum muricatiforme. In the by comparing it with that of other mountainous regions middle of the 20th century (Palamar-Mordvintseva 1978) of Europe – the Alps, Tatras and Sudetes. She noted 80 the dominants were Staurastrum senarium f. senarium taxa at species and intraspecific rank from six groups: and f. tatrica, Euastrum pinnatum, E. humerosum var. Cyanoprokaryota (3), Chrysophyta (1), Dinophyta (2), humerosum and var. affine, E. didelta, E. denticulatum Bacillariophyta (17), Chlorophyta (4) and Charophyta and Closterium directum. In the early 21st century the (43, according to the modern understanding of the rank dominants were Hyalotheca dissiliens, Netrium digi- and structure of the species as well as the identity of some tus, Euastrum humerosum var. affine, E. ansatum and of them). This information remained unchanged for almost Staurastrum polytrichum. The recently noted algal blooms half a century. Later, collections from 1967 (collected in Nesamovyte Lake, resulting from massive growth of by Z.I. Asaul) enabled regular study of the Desmidiales the green colonial coccoid alga Botryococcus terribilis, algae of the lakes of the Chornohora region, including confirm the instability of the lake’s ecosystem and its Nesamovyte and Brebeneskul lakes (Palamar-Mord- increase in trophy (Tsarenko et al. 2015, 2016, 2018; vintseva 1978a, b, 1982). However, these publications Mykitchak 2017). 56 Plant and Fungal Systematics 64(1): 53–64, 2019 ...... e m ot m m m m m m m m m ot m m me Тrо o-m o-m o-m o-m o-m o-m o-m o-m o-m o-m ...... SI 1.3 0.8 0.7 0.5 1.0 0.7 0.8 1.2 0.5 0.6 0.4 0.4 0.5 0.5 ...... o o o x-b o-x x-o o-x x-b x-o o-x x-o x-o x-o x-o Sap ...... i i hb hb hb Hal ...... acf acf acf acf acf рН acf acf acf acf acf acf acf acf acf acf acf acf acf acf acf ind ind ind ind ind ...... st ae ae Oxy st,ae S ...... B B B B B B B B B B B B B P-B P-B P-B P-B P-B P-B P-B Hab P, B B, . – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – + + 2013–2016 – – – – – – – – – – – – – – – – – – – – – – – – – – + + + + + + + + +

Brebenskul Lake 1967 – – – – – – – – – – – – – – – – – – – – 1 – – – – – – – + + + + + + + 2013-2016 – – – – – – – – – – – – – – – – – – – – – – – – – – + + + + + + + + + 1967 Nesamovyte Lake . – – – – – – – – – – – – – – – + + + + + + + + + + + + + + + + + + + 1920 Gutw. excavatum (Eichl. & & West G.S. Gutw.) West denticulatum papilliferum annulatum Eichl. & Gutw. humerosum parallelum (Willi Krieger) Kouwest & Coesel (Lund.) Teil. Gutw. angusticeps Groenbl. (West minus & (West G.S. Kurt West) Forster Menegh. ex Ralfs Eichl. & forma Gutw. tatrense Gutw. (West crassum & (West G.S. Grönblad West) tatrica Lütkem. Ralfs Ralfs ex Ralfs (Bréb.) W. Archer var. Archer C. venustum var. (Bréb.) W. C. umbilicatum Lütkem. C. staurastroides var. C. subcostatum var. C. staurastriforme Gutw. W. Archer C. pygmaeum W. C. polonicum Racib. C. obtusatum (Schmidle) Schmidle C. difficile C. margaritiferum C. minutum Delponte forma C. nasutum f. Actinotaenium clevei Actinotaenium A. cucurbita (Bréb. ex Ralfs) ex Teiling Růžička Таxa var. C. costatum var. brebissonii (Ralfs) Cylindrocystis de Bary var. A. silvae-nigrae var. C. brebissonii Menegh. ex Ralfs Euastrum abоense Elfv. Closterium archerianum Cleve Lundell ex P. C. intermedium Ralfs C. bipunctatum Børgesen E. ansatum (Ehenb Ralfs.) C. rostratum Ehrenb. ex Ralfs C. navicula var. C. striolatum Ehrenb. C. angulosum Bréb. C. jenneri Cosmarium amoenum var. E. binale (Turp.) Ralfs var. E. didelta E. denticulatum (Kirchn.) Gay var. E. elegans (Bréb.) Kütz. var. E. denticulatum var. E. humerosum Ralfs var. Table 2. The algal indicators Table of the Nesamovyte and Brebenskul Lakes with species ecology (Barinova et al., 2006, 2019). Habitat preferences: B – benthic; P-B – planktic-benthic; S – soil; Ep – epiphytes; – P planktonic. аer – aerophyles. Streaming and oxygenation: St-str, low-flow moderately oxygenated water inhabitants; ae – aerophytes;– stacidophils. – Salinity: standingi water– indifferents; with hb low – oxygenation halophobes; inhabitants. oh – pH olihohalobes.Trophic (Hustedt state 1957): Dam (Tro) (Van ind et – al. 1994): indifferents; m acf oligotraphentic. –pollution Organic (SI, Sap): x mesotraphentic;– xenosaprobes, x-o – mexeno-oligosaprobes, o-x –oligo-xenosaprobes, x-b –– xeno-betamesosaprobes. o – meso-eutraphentic;oligosaprobes, o-b – o-moligo-betamesosaprobes, b-o – beta-oligosaprobes, – oligo- to mesotraphentic; e – eutraphentic; ot – o-a – oligo-alphamesosaprobes, b – betamesosaprobes, b-a – beta-alphamesosaprobes, a – alpha-mesosaprobes. P. Tsarenko et al.: Green and charophytic algae of the high-mountain lakes in the Eastern Carpathians 57 ...... m ot m ot m m ot ot m m m m ot m m m m Тrо o-m o-m o-m o-m o-m o-m o-m o-m o-m o-m ...... SI 1.2 1.5 1.6 0.5 0.5 0.5 0.5 0.5 0.5 0.6 0.8 0.9 0.5 0.3 0.4 0.5 0.5 0.4 0.9 0.5 ...... o x o-b b-o x-o x-o x-o x-o x-o x-o o-x x-b x-b x-o x-o x-o x-o x-o x-b x-o Sap ...... i i i i hb hb hb Hal ...... acf acf acf acf acf acf acf acf рН acf acf acf acf acf acf acf acf acf acf acf acf acf acf ind ind ind ind ind ...... st ae ae ae Oxy st-str st-str st-str ...... P P B B B B B B B B B B B B B P-B P-B P-B P-B P-B P-B P-B P-B Hab B, aer B, aer B, aer – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – + + + + + + 2013–2016 . – – – – – – – – – – – – – – – – – – – – – – – – – – – + + + + + + + + + + Brebenskul Lake 1967 – – 1 2 – – – – – – – – – – – – – – – 2 2 – – – – – – – – – + + + + + + + + 2013-2016 . – – – – – – – – – – – – – – – – – – – – – – – + + + + + + + + + + + + + + 1967 Nesamovyte Lake . . – – – – – – – – – – – – – – – – – – – – – – + + + + + + + + + + + + + + 1920 Nordst. (West) Coesel (West) & Meesters

pilosellum muticum West & West G.S. West subavicula (Ralfs) Racib. var. obesum affine subintermedium Schröd. austriacum Lütkem. West cylindricum & West G.S. West muricatum (Bréb. ex Ralfs) Kurt Förster tatrica Gutw. subangulare var. sp.st Bréb. S. hexacerum Wittrock S. pilosum Bréb. S. polytrichum (Perty) Rabenh. S. diacanthum Lemaire S. dilatatum Ehrenb. ex Ralfs var. S. hirsutum var. S. muricatiforme Schmidle S. muricatum f. S. punctulatum Bréb. E. humerosum E. insigne Hassall ex Ralfs West S. conspicuum & West G.S.West S. muticum Bréb. ex Ralfs var. var. E. humerosum var. Таxa West E. montanum & West G.S. West E. pinnatum Ralfs E. obesum Josh. var. var. E. obesum var. E. tuddalense Ström E. verrucosum Ehrenb. ex Ralfs Netrium digitus (Ehrenb. ex Ralfs) Itzigs. & Rothe emend. Ohtani var. Gonatozygon monotaenium var. Hyalotheca dissiliens Hyalotheca Bréb. ex Ralfs H. mucosa (Mertens) Ehrenb. Penium cylindrus Bréb. ex Ralfs Micrasterias fimbriata Ralfs Mougeotia N. oblongum (De Bary) Lütkm. var. N. oblongum var. spirostriolatum J. Barker P. S. coarctatus Breb. Spirotaenia condensata Spirotaenia Bréb. ex Ralfs S. avicula (Ralfs) ehrenbergii De Pleurotaenium Bary Sphaerozosma aubertianum West Sphaerozosma Spirogyra sp. var. S. pulchellum var. Staurastrum arcuatum Table 2 . Continued. Table 58 Plant and Fungal Systematics 64(1): 53–64, 2019 ...... ot m m m m m m m Тrо o-m o-m o-m o-m o-m o-m ...... SI 1.8 2.1 2.1 1.3 1.5 0.8 3.1 2.3 0.8 1.5 0.5 2.15 ...... a b b o b b o-a o-b x-b x-b o-b x-o Sap ...... i i i i i hb oh Hal ...... acf acf acf acf acf acf acf acf acf acf acf рН ind ind ind ind ind ind ind ind ...... st ae ae Oxy st-str st-str st-str st-str st-str st-str st-str ...... P P P B B B B B B B B P-B P-B P-B P-B P-B P-B Hab B, aer B, aer P-B, Ep . . – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – + + + + 2013–2016 – – . . – – – – – – – – – – – – – – – – – – – – – – – – – – – – – + + + + +

Brebenskul Lake 1967 . – – – – – – – – 5 – – – – – – – – – – – – + + + + + + + + + + + + + + + + + 2013-2016 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – + + + + + + + + 1967 Nesamovyte Lake . . – – – – – – – – – – – – – – – – – – – – – + + + + + + + + + + + + + + + + 1920 Hegew. var. boryanum var. Hegew. Bock & Krienitz Hegew. brebissonii Dangeard Racib. senarium (Nauman) Komárek & Perman (H.C. C. Wood) Bock, Pröschold & Krienitz Hegew.) E. Hegew.) P.A. tatrica var. levis var. (Bréb. ex Ralfs) Teiling teliferum (Roy & Bisset) Bourrelly (West) West & West G.S. productum (West) West minor De Bary tyrolense (Schmidle) Irenne-Maria (Reinsch) P. longicorne Tsarenko (Reinsch) P. (W.B. Turner) Turner) gladiosum Coesel (W.B. & Meesters isthmosus (Heimerl) Coesel (Woronich.) intermedius Růžička (Woronich.) Ralfs var. Reinsch granulata var. var. (West) botryoides De (West) Wildeman Westella Tetradesmus dimorphus (Turpin) M.J. Tetradesmus Wynne var. boryanum var. P. Pseudopediastrum boryanum (Turpin) E. P. duplex Meyen P. Pediastrum braunii Wartmann Oedogonium sp. st. var. S. extensus var. Staurodesmus dejectus Staurodesmus brebissonii Ralfs var. Tetmemorus var. brebissonii var. T. Botryococcus braunii Kütz Botryococcus Chlamydomonas reinhardtii Chlamydomonas reinhardtii Coelastrum sp. Mychonastes anomalus (Korschikov) Krienitz, C.Bock, Dadheech & Pröschold S. vastum Schmidle S. glaber (Ehrenb. ex Ralfs) Teiling Zygnema sp. st. B. terribilis Komarek B. terribilis & Marvan Mucidosphaerium pulchellum var. var. S. teliferum S. turgescens De Not. S. quadratus (Schmidle) Teiling Teilingia T. laevis Ralfs T. ex Ralfs Tortitaenia alpina (Schmidle) Tortitaenia Brook Lemmermannia triangularis (Chodat) C. S. sexcostatum Bréb. ex Ralfs var. S. sexcostatum var. S. scabrum Bréb. S. teliferum T. laevis T. Staurastrum senarium f. var. S. subavicula var. Desmodesmus communis (E. S. senarium (Ehrenb.) Ralfs f. S. sebaldii Dictyosphaerium chlorelloides Таxa (Wille) P.Tsarenko & D.M. John irregularis Crucigeniella (Wille) P.Tsarenko Table 2 . Continued. Table P. Tsarenko et al.: Green and charophytic algae of the high-mountain lakes in the Eastern Carpathians 59

Nesamovyte Brebeneskul of mineralization, sharp temperature gradient from the upper to bottom layers of the water, and the lake’s ori- gin and location (at the bottom of a glacial cirque). The presence of rare or typical species (Gonatozygon mono-

86 8 18 taenium var. pilosellum, Netrium digitus, Staurastrum (76.8%) (7.1%) (16.1%) diacanthum, Staurastrum punctulatum) testifies to the algosozological value of this lake, the appropriateness of its being included in the Carpathian Biosphere Reserve, and the need for environmental protection measures to maintain this ecosystem. Figure 2. Degree of similarity of species composition of green and The two lakes differed in algal species composition. charophytic algae between Nesamovyte and Brebeneskul lakes. Most of the taxa (83 species, 92 intraspecific; ~84%) were recorded in Nesamovyte Lake. The degree of sim- The algae of Brebeneskul Lake show low floristic and ilarity between the green and charophytic algae of this taxonomic diversity, with only 23 species (26 intraspe- lake and those of Brebeneskul Lake was very low, only cific), which is ~24% of the total algal species of the 7.1% (Fig. 2). investigated lakes. Charophytes account for ~77% of the Only seven taxa were found in the material from both identified algae; green algae are less diverse (23%) and lakes. (Cylindrocystis brebissonii, Euastrum denticulatum, are found occasionally, mostly in the marshy part of the Euastrum denticulatum var. angusticeps, Netrium lake environment among higher aquatic plant associations. digitus, Penium cylindrus, Desmodesmus communis, The species composition of the plankton and benthos (the Pseudopediastrum boryanum var. longicorne). The dif- rocky bottom of the lake, slightly silted near shore) was ferences in the algal species composition of the two lakes depauperate. The most diverse genera are Staurastrum, can be explained by environmental factors and by fea- Cosmarium, Euastrum, Closterium and Pseudopedias- tures specific to these waterbodies – differences in their trum, and the remaining eight genera of both groups are morphometric indicators, and the degree of modern trans- represented by one species each. Such low species diver- formation of their ecosystems. In particular, the species sity in Brebeneskul Lake was noted earlier for eugleno- diversity of Nesamovyte Lake apparently has decreased phytes and desmids (Аsaul 1969; Palamar-Mordvintseva as a result of anthropopression, changes in trophy, tem- 1978а); this was apparently due to its oligotrophy, degree perature increases and long dry periods in recent years,

A 14 16 C III I B II 12 12 14 10

8 10 8 6 8 6 4 6 4 No. of No. of species No. of No. of species No. of No. of species 2 4 2 2 0 0 P P-B P-B, Ep B B, aer B,S 0 P P-B P-B, Ep B B, aer P,B P-B B B, aer Ecological category Ecological category Ecological category

6 2.5 7 III D I E F II 6 5 2

5 4 1.5 4 3 1 3 2 2 No. of No. of species No. of No. of species No. of No. of species 1 0.5 1

0 0 0 st-str st,ae ae st ae st st-str ae Ecological category Ecological category Ecological category 30 I H 20 II I III G 18 16 25 16 14

20 14 12 12 10 15 10 8 8 6 10 6 No. of No. of species No. of No. of species No. of No. of species 4 5 4 2 2 0 0 0 acf ind acf ind acf ind Ecological category Ecological category Ecological category

Figure 3. Bioindication plots for ecological analysis of habitat preference (A–C), current and oxygenation (D–F) and pH (G–I) in Nesamovyte Lake (for the periods: I – 1920, II – 1967, III – 2013–2016). 60 Plant and Fungal Systematics 64(1): 53–64, 2019

4 A I B 2 II C 6 III 5

3 1.5

2 1 3 2 No. of species No. of species

0.5 No. of species 1 1 0 0 0 hb hb i oh hb i Ecological category Ecological category Ecological category

D I E II F III 14 10 18 9 12 8

14 10 7 12 6 10 8 5 8 6 4 3 No. of species

6 No. of species 4 4 No. of species 2 2 2 1 0 0 0 ot o-m m ot o-m m ot o-m m Ecological category Ecological category Ecological category G 14 H 12 I 7 III I II 12 10 6

10 5

8 8 4 6 6 3 4

No. of species 2

4 No. of species

2 No. of species 2 1 0 0 0 I II III I II III I II III IV Water quality Class Water quality Class Water quality Class

Figure 4. Bioindication plots for ecological analysis of salinity (A–C), Trophic state (D–F), water quality class (according to organic pollution) (G–I) in Nesamovyte Lake (for the periods: I – 1920, II – 1967, III – 2013–2016).

resulting in a decrease of the lake surface area and lower was a more or less uniform distribution of taxa indi- water content of the near-shore wetland, and the lack of cating low-flow, moderately oxygenated water (st-str) characteristic types of oligotrophic waterbodies in the and standing water with low oxygenation (st) (Fig. 3D). subalpine zone (Wołoszynska 1920; Palamar-Mordvint- Data from 1967 show a gradual increase in the role of seva 1978a, 1982; Lukavsky 1994; Lenarczyk & Tsarenko standing-water algae, with taxa indicating low oxygen- 2013). Similar changes in the ecosystem and in its species ation. By 2013–2016, the dominant taxa had shifted from composition of algae have also occurred in Brebeneskul aerophytes to taxa characteristic of low-flow, moderately Lake. oxygenated water (st-str) (Fig. 3E, F). This indication of The changes in algal indicator taxa algae in Nesam- lower oxygen may reflect environmental degradation. ovyte Lake in 1920, 1967 and 2013–2016 reflect changes The pH-indicating taxonomic groups remained sta- in lake conditions. Taxa that inhabit different biotopes ble over the studied years: only indifferents (ind) and declined. In 1920 the dominant indicators were benthic acidophiles (acf) were found. Acidophiles prevailed (B) along with plankto-benthic (P-B), followed by ben- in all the studied years, indicating neutral to slightly thic and aerophyles (B-aer), planktonic (P), benthic and acid water (Fig. 3G, H, I), but the increasing share of soil (B, S), and plankto-benthic and epiphytes (P-B, Ep); indifferents suggests an upward trend of pH from neutral in 2014–2016 we noted decreases of P-B, B, B-aer, P to 7, which is seen in Nesamovyte Lake (2013–2016) and P-B, and Ep, along with a decrease in the number (Fig. 3I). In terms of salinity, indicators of typically fresh of benthic forms, the predominance of plankto-benthic waterbodies were identified, characterizing the lakes as and disappearance of a group of algae that can occur in freshwater both earlier and recently, though we observed both benthos and soil (Fig. 3A, B, C). Perhaps this can a tendency to increased mineralization, the appearance be explained by the changes in the hydrological regime of low numbers of oligohalobes (Fig. 4A, B, C). The or by differences in the way algological material was trophic state of the lake is determined by a multitude of sampled in the 1920s, as well as by the current state of interrelated physicochemical and biological processes; in the waterbody and its near-shore wetland. these lakes those processes were reflected in the presence Data on the algal indicators of oxygen regime and of mesotraphentic (m), oligo- to mesotraphentic (o-m) and water movement confirm changes in the hydrological oligotraphentic (ot) (Fig. 4D, E, F) indicator groups. In the regime of the lake. Early data indicate the predominance recent period, as before, oligo- to mesotraphentic indica- of aerophytes (ae), indicating that the water was oxy- tors prevailed, reflecting relatively clean water. However, gen-enriched in 1920; along with the aerophytes there as seen from Figure 6, a decrease of oligotraphentic P. Tsarenko et al.: Green and charophytic algae of the high-mountain lakes in the Eastern Carpathians 61

A 9 I B 6 II 8 5 7

6 4 5 4 3 3 2 No. of No. of species 2 No. of species 1 1 0 0 P P-B P-B, Ep B B,S P P-B P-B, Ep B Ecological category Ecological category

C 8 I D 6 II 7 5

5 4 4 3 3

No. of No. of species 2

No. of No. of species 2 1 1 0 st-str st,ae 0 st-1str Ecological category Ecological category

E 14 I F 4.5 II 4 12

3.5 10 3 8 2.5 6 2 1.5 No. of No. of species 4 No. of species 1 2 0.5 0 0 acf ind acf ind Ecological category Ecological category

Figure 5. Bioindication plots for ecological analysis of habitat preference (A, B), current and oxygenation (C, D) and pH (E, F) in Brebeneskul Lake (for the periods: I – 1967, II – 2013–2016). indicators and the growing share of mesotraphentic spe- aerophytes (ae), were in the 1967 samples but not in the cies testify to deterioration of lake trophy. In regard to material from 2013–2016. This supports the suggestion organic pollution, taxa indicating the first (cleanest) class of altered hydrology, but in view of the low number of of water quality prevail (Fig. 4G, H, I). In Nesamovyte indicators it is difficult to assert that with certainty. In Lake the indicator taxa were distributed among the three Brebeneskul Lake, in both 1967 and 2013–2016 the same water quality classes (I, II, III) at the beginning of the groups of water pH indicators were found, but in 1967 20th century (1920) and also in subsequent periods (1967, acidophiles dominated and there were few indifferents; 2013–2016). Recently, however, taxa indicating quality the share of indifferents has increased since then, point- class V class were found; no taxa of that group were ing to an increase in pH, closer to pH 7 (Fig. 5E, F). present in 1920 and 1967. Thus we see some deterioration For salinity, indiferrents predominated both recently and of the state of the lake, despite the continued dominance earlier, but halophobes (hb), present in 1967, no longer of taxa indicating water quality class I. occur in the lake (Fig. 6A, B). Halophobes are strictly In the ecosystem of Brebeneskul Lake, plankto-ben- freshwater species, disappearing with a slight increase in thic species dominated the algal assemblage in both salinity. Thus, it can be assumed that small changes in the 1967 and 2013–2016. The shares of other habitat-lim- mineral regime have occurred, though the water remains ited organisms changed over time. In 1967 the number fresh. Figure 6C, d shows the decline in the trophic state of of benthic forms was higher; it decreased in 2013–2016, the lake through time. In the earlier period, mesotraphentic replaced by a larger number of planktonic forms (Fig. indicators prevailed, followed by oligo- to mesotraphentic 5A, B); differences in sampling methods may be partly (o-m); eutraphentic (e) and oligotraphentic (ot) indicators responsible for this. The 1967 samples contained benthic, were marginally represented. The picture has changed plankto-benthic, planktonic, benthic, soil, plankto-benthic somewhat since then: oligo- to mesotraphentic species and epiphytic taxa; by 2013–2016 the benthic and soil (o-m) are followed by mesotraphentic and a small number species had vanished. This suggests a change in the lake’s of meso-eutraphentic (me). hydrological regime. Here we note that taxa of low-flow, Organic pollution gives the final evidence about moderately oxygenated water were observed in the recent changes in the state of Brebeneskul Lake (Fig. 6E, F). period as well as earlier (Fig. 5C, D). Taxa indicating In 1967, indicators of water quality class II prevailed, standing water (st) with low oxygenation, along with with few indicators of classes I and III; the material from 62 Plant and Fungal Systematics 64(1): 53–64, 2019

B A 8 I 6 II 7 5 6

5 4 4 3 3

No. of No. of species 2

2 No. of species 1 1 0 0 hb i 1 i Ecological category Ecological category

C 9 I D 3.5 II 8 3 7

2.5 6 5 2 4 1.5 3

No. of No. of species 1 No. of No. of species 2 0.5 1 0 0 ot o-m m e o-m m me Ecological category Ecological category 7 E I F 3.5 II 6 3

5 2.5 4 2 3 1.5

No. of No. of species 2

No. of No. of species 1 1 0.5 0 0 I II III I II III Water quality Class Water quality Class Figure 6. Bioindication plots for salinity (A, B), trophic state (C, D) and water quality class (according to organic pollution) (E, F) in Brebeneskul Lake (I: 1967, II: 2013–2016).

2013–2016 contains the same shares of class III and class deteriorated recently, and that the negative changes are II indicator taxa as noted in 1967, and fewer class I indi- worse in Brebeneskul Lake. cators. From this we infer that the water has changed from clean to moderately polluted. Acknowledgements

We thank Professor I.I. Chorney, Associate Professors V.V. Conclusions Budjak and O.I. Khudyi (Yuri Fedkovych Chernivtsi University) Our bioindication-based environmental analysis focus- and Dr. T. Mykytchak (Institute of Ecology of the Carpathians, ing on the composition of green and charophyte algae National Academy of Science of Ukraine, Lviv) for organizing and conducting fi eld trips to the Ukrainian Carpathians and produced conclusions restricted to several systematic for assistance in the work, and Dr. J. Tunovsky (Institute of groups. In terms of typology and biology, the two studied Freshwater Fish Research, Poland) for the providing data on high-mountain lakes of Chornohora are oligo-mesotrophic hydroecological indicators. Financial and organizational support and contain the corresponding range of aquatic organisms was provided by the administration and staff of the National and algae. However, increased anthropopression, impacts Academy of Science of Ukraine and the Polish Academy of of recreational use, and the accompanying changes in the Sciences for support of Polish–Ukrainian scientifi c cooperation. ecosystem are transforming their biota. Such changes characterize the Nesamovyte Lake ecosystem over the References more than century-long observation period: in particular, the transition from oligotrophic to beta-mesosaprobic Andersen, R. A. (ed.). 2005. Algal Culturing Techniques. Elsevier Acad. (Wołoszynska 1920; Аsaul 1969, Palamar-Mordvint- Press, London. seva 1978a, b, 1982; Mykitchak 2014). Confi rmation is Asaul, Z. I. 1969. Euglenophyta of high mountain lakes of the Ukrainian found in the results of studies of lake hydrobionts from Carpathians. Ukrainian Botanical Journal 26: 8–13. 2012–2013 (Mykitchak et al. 2014; Tsarenko et al. 2014, Bąk, M., Witkowski A., Żelazna-Wieczorek J., Wojtal A. Z., Szcze- Mykitchak 2017). Our study of the green and charophyte pocka E., Szulc A. & Szulc B. 2012. Klucz do oznaczania okrzemek w fi tobentosie na potrzeby oceny stanu ekologicznego wód powierzch- indicator species of the two lakes suggests that the ecosys- niowych w Polsce. Warszawa: Główny Inspektorat Ochrony Śro- tems of Nesamovyte Lake and Brebeneskul Lake have dowiska. P. Tsarenko et al.: Green and charophytic algae of the high-mountain lakes in the Eastern Carpathians 63

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